ActaM. Geraldes Botanica et alMalacitana. Genista in 39. Iberian 45-54 Peninsula and Morocco Málaga, 201445 NEW GENETIC DATA ON GENISTA ANGLICA L. VERSUS GENISTA ANCISTROCARPA SPACH (FABACEAE, FABALES) IN THE IBERIAN PENINSULA AND MOROCCO. PHYLOGEOGRAPHIC CLUES Miguel GERALDES1*, João Paulo FONSECA2, Carlos NETO1 & José Carlos COSTA2 1 Centro de Estudos Geográficos, Instituto de Geografia e Ordenamento do Território, Universidade de Lisboa, Alameda da Universidade 1600-214. Lisboa. Portugal 2 Centro de Biociências. ISPA-IU. R. Jardim do Tabaco, 34, 1149 - 041 Lisboa. Portugal 3 CBAA, Instituto Superior de Agronomia, Universidade de Lisboa, Tapada da Ajuda, 1349017 Lisboa. Portugal * Correspondence author: [email protected] Recibido el 20 de enero de 2014, aceptado para su publicación el 31 de enero de 2014 ABSTRACT. New genetic data on Genista anglica L. versus Genista ancistrocarpa Spach (Fabaceae, Fabales) in the Iberian Peninsula and Morocco. Phylogeographic clues. The genetic analysis through the nuclear ribosomal DNA of some Iberian and NW Morocco populations of Genista ancistrocarpa Spach and G. anglica L. has validated the separation of these two sister taxa as two distinct genetic entities, strengthening the already described morphological differentiation. The studied populations neatly show the G. ancistrocarpa as a coastal taxon with occidental Iberian and NW Morocco distribution, contrary to the G. anglica. The distribution areas of the two species appear to be distinct and with an important biogeographical significance. The coastal clade is determined by populations of G. ancistrocarpa, and outlines a biogeographic region that seems to depart genetically from the remaining peninsular populations, also in several molecular analyses of other plants and animals by many authors, supporting the idea of a distinct evolution from the Miocene onwards, when the installation of dry and hot conditions has pushed the moisture-dependent populations of G. ancistracarpa toward the westernmost sectors of Iberia and Morocco with an atlantic influence. Key words. Chorology, Genista anglica, Genista ancistrocarpa, sublittoral fens, Iberian Peninsula, Morocco. RESUMO. Novos dados genéticos sobre a Genista anglica L. versus Genista ancistrocarpa Spach (Fabaceae, Fabales) na Península Ibérica e Marrocos. Indícios filogeográficos.A análise genética de ADN nuclear ribossomal, levada a cabo em algumas populações peninsulares e do NW de Marrocos de Genista ancistrocarpa e de G. anglica, permitiu validar estes dois taxa irmãos como entidades genéticas distintas, reforçando, desta forma, a sua diferenciação morfológica. As análises permitem definir aG. ancistrocarpa como um taxon litoral com distribuição oeste ibérica e NW de África, por oposição à G. anglica. As áreas de distribuição das duas espécies parecem ser distintas e com importante significado biogeográfico. O This work was supported by the Portuguese Foundation for Science and Technology (FCT) through the project FCT - PTDC/AAC-AMB/111349/2009. 46 Acta Botanica Malacitana 39. 2014 clado litoral definido pelas populações de G. ancistrocarpa define um território biogeográfico, que parece destacarse geneticamente das restantes populações peninsulares, em diversas plantas e animais analisados geneticamente por vários autores, reforçando a ideia de uma evolução distinta a partir do Mioceno médio, quando a instalação de um clima seco e quente empurrou e isolou uma parte da população da higrófila G. ancistrocarpa para ocidente, onde podiam receber uma influência atlântica. Palavras-chave. Corologia, Genista anglica, Genista ancistrocarpa, turfeiras sublitorais, Península Ibérica, Marrocos. INTRODUCTION and new data from nuclear ribosomal DNA for these very closely-related taxa. While the most species of the Genista It has indeed been accepted that G. anglica genus thrive on dry soils, the Genista anglica and G. ancistrocarpa are morphologically L. and Genista ancistrocarpa Spach are two analogous in many phenotypic traits. In species of atlantic kinship, which have their the Iberian Flora (Talavera, 1999), they are southernmost chorology on wet, acidic, poor, separated mainly by the number of leaflets in the peat-rich soils in Portugal, Spain and Morocco, armpit of the infertile talus, being unifoliated commonly near drainage lines with low energy in G. anglica, yet trifoliated in non-flowering like creeks, ponds and lakes, whether very near branches in G. ancistrocarpa. to the coastline (sublittoral ecosystems) or in In the Iberian Peninsula, the G. the non-arid mountains (Costa et al. 1998). ancistrocarpa occurs mainly in lowlands These habitats occur where annual mean (0-20 m), near the ocean, with local edaphic rainfall reaches over 800 mm up to 2800 mm fresh-water compensation, originating peaty or, concerning areas more to the south, where fen complexes neighboured by hygrophilous the water-table lies near the surface and springs heathlands, while G. anglica appears at higher out, creating water-logged conditions. elevations (10-1800 m), where greater values Recent taxonomic discussion has been of annual precipitation are recorded, normally performed about the other southern chorology, more than 800 mm (Talavera, 1999). the isolated populations in Calabria, southern Italy, on two of its mountain ranges - the Study area Aspromonte and the Sila, formerly considered In order to clarify the phylogenetic as G. anglica, which have been recently relationships among populations of the two described as a new endemic taxon, G. silana species (G. ancistrocarpa and G. anglica) Brullo, Gangale & Spamp for Sila range and of the Southwestern Iberian Peninsula and G. brutia Brullo, Scelsi & Spamp. (Brullo et al. Northwestern Morocco, we performed some 2001) for the Aspromonte range, albeit not yet genetic analysis using specimens of G. anglica acknowledged by a great many authors. These sampled in the following places: Sierra de will not be considered in this article. Cebollera (Northern Iberian System Range), The present work aims to contribute to Sierra de los Ancares (Cantabrian System shed light on the systemic reports between Range), Serra da Estrela (Central Iberian southwestern populations of the two commonly System Range), and Morais Massif (Trás-os- accepted sister taxa of wet–soil Genista: G. Montes). For the G. ancistrocarpa, the leaves anglica and G. ancistrocarpa, exploring the were sampled from the Graben of Chaves comparison between pheno-taxonomic traits (Trás-os-Montes), the Serra de Montemuro M. Geraldes et al. Genista in Iberian Peninsula and Morocco 47 (Northwestern Portuguese Mountains), the Sado Estuary (Alentejo Coast), and Jbel Bou Hachem (Western Rif Range), as we can observe in figure 1. In the southwest of the Iberian Peninsula, the lowland specimens that we sampled, mainly from the Setúbal Peninsula, Sado Estuary and Alentejo Coast (Portugal), Doñana (Spain), and, over the Strait, in Larache and the Rif (Morocco), exhibited morphological characteristics that enabled the identification as G. ancistrocarpa. The specimens that we have observed at two sites on the Western Rif Mountains in Morocco (in Bou Hassim region, above Boubiyene, 1019 m, and above El Maouzkir, 1200 m) clearly showed all trifoliate leaves, hence being, after the Iberian Flora key, also G. ancistrocarpa. According to many authors, the specimens Figure 1. Location of the sampled populations of G. from those regions were identified as Genista ancistrocarpa and G. anglica. Genista ancistrocarpa:1- Veiga de Chaves - ancistrocarpa (Azzioui et al. 2000; Benito et 41°45’29,24”N; 7°27’13,34”W; 350 m. 2- Serra de al. 2006; Romo, 2009; Chambouleyron, 2012) Montemuro - 40°58’13,74”N; 7°59’20,84”W; 1352m. or as the synonym Genista anglica subsp. 3- Sado Estuary - 38°23’04,91”N; 8°36’04,27”W; 25 m. 4- Western Rif (Jbel Bou Hachem) - ancistrocarpa Maire (Deil et al. 2010); such 36°15’33,45”N; 5°26’30,50”W; 1019m. Genista information is available in the Anthos online anglica: a- Sierra de Cebollera - 42°07’00,63”N; database also (Anthos, 2013). In both cases, the 2°34’09,63”W; 1100m. b- Sierra de Los Ancares - 42°51’38,20”N; 6°51’20,70”W; 1100m. c- Morais sampled specimina occurred on tiny swampy Massif - 41°31’33,52”N; 6°48’17,34”W; 650 m. depressions with a confined low-rate runoff d- Serra da Estrela - 40°21’33,62”N; 7°39’09,06”W; of groundwater. The surface water had only 1584 m. a few inches depth in the Summer, while it oscillates significantly during the year due to the height where the latter can occur. The Western Mediterranean climate, under which an aestival Rif Mountains represent the areas with the absence of rainfall prevails. In the Winter, we highest rainfall records in Morocco, with values have observed a small rise, but the runoff was always over 600 mm, reaching even 2100 mm slow as well. in some of its highest summits (Rouai & Jaaidi, Using the ecological distribution model 2003). The location of the Rif Mountains, from of these two species in the Iberian Peninsula, where the specimens from G. ancistrocarpa the rainfall and height that characterize both were collected (available on the herbarium sites in the Rif Mountains would predict them João Carvalho & Vasconcellos – code LISI), to be included within the potential range of G. have the highest precipitation values in anglica, not the ‘lowland’ G. ancistrocarpa. the whole Rif Chain and hence across the However, the morphological and genetic northwest of Morocco. This climatic originality data indicate that the sampled plants were individualizes the Western Rif as one of the 20 unmistakably G. ancistrocarpa, despite the natural areas in which the North of Morocco is elevation, increasing thereby substantially the divided (Valdés, 2006; Molina-Venegas et al. 48 Acta Botanica Malacitana 39. 2014 2013). It is also the natural area with the highest 1999), and also the Clustal X software, version floristic diversity, with an impressive total of 1.81 (Thompson et al. 1997). For the initial 1758 taxa. The occurrence of G. ancistrocarpa approach to the phylogenetic analysis, we used at high rainy areas, such as the Western Rif the Bayesian inference through the Mr.